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Ernst  Mayr  Library 

of  the  Museum  of 

Comparative  Zoology 







Zoological  Institute 

Academy  of  Sciences 

Leningrad ,  USSR 



Transactions  of  the  Zoological  Institute 
Academy  of  Sciences,  USSR,  Vol.  80,  pp.  kh-69 ,   1979 

An  edited  translation  of: 

Sistema  Otryada  Geophila  {=Helicida)  (Gastropoda  Pxilmonata) 
Morfologiya,  Sistematika  i  Filogeniya  Mollyuskov 
Trudy  Zoologicheskogo  Instituta,  Akademiya  Nauk  SSSR 
Tom  80,  pp.  l+U-69,  1979 


Museum  of  Comparative  Zoology 
Harvard  University 
Cambridge,  Ma.   02138 

Special  Occasional  Publication 

NO.   6 

published  by  the 

Department  of  Mollusks 

Heirvard  University 

Cambridge,  Massachusetts  02136 



A.  A.  Schileyko 

Generally  speaking  the  fewer  the  characteristics  employed  as 
bases  for  the  systematics  of  any  group  of  animals,  the  more  consis- 
tently harmonious  that  taxonomic  scheme  seems  to  be.   When  using  numerous 
characteristics  relating  to  particular  organs  or  structures,  one  often 
elaborates  a  less  harmonious,  though  more  synthetic,  system  because 
certain  groups  of  features  fail  to  correspond  to  others.   Also  the 
more  adequate  system  is  strengthened  by  the  utilization  of  a  holistic 
approach  wherein  the  various  morphological  systems  are  viewed  as  inter- 
related parts  of  an  organic  whole,  the  result  of  the  dynamic  forces 
of  evolution.   At  higher  taxonomic  ranks,  the  indicators  of  evolutionary 
and  developmental  pathways  of  the  various  organs  and  structures  take 
on  greater  significance. 

T\irning  our  attention  to  the  current  systematics  of  the  Order 
Geophila,  we  recognize  that  attempts  to  understand  their  phyletic 
relationships  have  fallen  short  of  a  holistic  analysis.   Reduction  of, 
and  substitution  for,  one  organ  by  another  has  been  frequent  while 
problems  of  parallelism  and  convergence  have  often  been  insufficiently 
appreciated.   Only  through  a  more  thorough  analysis  of  such  phenomena, 
including  a  study  of  variation  between  species  and  among  genera  as 
well  as  the  discernment  of  evolutionary  pathways  will  a  more  com^pre- 
hensive,  phyletically  accurate  system  emerge. 

Various  opinions  of  the  systematics  of  the  order  exist.   Most 
specialists  accept  the  one  proposed  by  Pilsbry  (l900)  which  is  based 

on  the  characteristics  of  the  structTxre  of  the  excretory  apparatus;  and 
he  divides  the  order  into  3  suborders:   Orthiirethra ,  Heterurethra, 
and  Sigmurethra.   Later  a  fourth  group,  Mesurethra,  was  added  by  Boettger 
(195U)  who  shared  Pilsbry's  opinions.   Solem  (1959)  proposed  a  phylogenetic 
system  based  on  the  unconnected  branches  Orthurethra,  Sigmurethra,  Mesurethra, 
and  Tracheopulmonata  plus  Heterurethra.   Thiele's  system  (l93l),  based 
primarily  on  conchological  features,  has  often  been  subjected  to  criticism 
from  the  point  of  view  of  the  anatomist,  although  the  validity  of  many  of 
Thiele's  formerly  rejected  arguments  should  now  be  recognized.   The  system 
proposed  by  Wenz  &  Zilch  (Zilch,  1959-196o)  is  a  compilation  in  which 
Thiele's  system,  modified  to  conform  to  newly  accumiilated  facts  (between 
193i+  and  1959),  was  superimposed  on  Pilsbry's  earlier  scheme.   The  most 
widely  currently  accepted  arrangement  is  the  one  proposed  by  Taylor  & 
Sohl  (1962),  where  the  same  suborders  appear,  divided  into  19  superfamilies. 

Moreover,  the  largest  group,  Sigmurethra,  was  divided  by  H.B.  Baker, 
following  Pilsbry,  into  the  infraorders ,  Holopoda  and  Aulacopoda,  plus  a 
new  infraorder,  Holopodopes.   The  members  of  these  groups  are  distinguished 
by  the  nature  of  furrows  or  grooves  in  the  foot.   In  Holopoda  and  Holopodopes 

Heterurethra  as  understood  by  Pilsbry  includes  3  independent  groups 
(accorded  ordinal  rank  by  Soviet  malacologists) :   Succineida,  Aylliida, 
Athoracophorida  (Minichev,  1971;  Minichev  &  Starobogatov,  1975;  Slavoshevskaya, 
197I;  Van  Mol,  I967).   For  this  reason  the  early  groups  of  Heterurethra 
will  not  be  treated  here. 

the  peripedal  fiirrow  runs  along  the  margin  of  the  sole  or  slightly  above 
it,  the  sole  itself  always  being  entire.   In  Aulacopoda  the  peripedal  furrow 
is  placed  noticeably  higher  than  the  margin  of  the  sole  and  the  sole  itself 
is  often  divided  by  2  parallel  furrows.   The  obstacle  to  uniting  Holopodopes 
and  Holopoda  lies  in  the  fact  that  Baker  assigns  them  to  different  phylo- 
genetic  branches:   the  first  is  lower  than  Sigmurethra,  the  second  is  higher. 

Hence  up  to  the  present,  three  groups  of  characteristics  have  been 
proposed  as  suitable  bases  for  the  construction  of  a  system  for  the  order 
Geophila:  conchological ,  excretory,  and  pedal.  In  addition,  the  sexual 
apparatus  has  been  used  in  diagnosis  at  the  familial  or  generic  levels. 
We  plan  to  examine  separately  a  series  of  groups  of  characteristics;  to 
simplify  matters,  we  will  limit  ourselves  to  the  discussion  only  of  the 
system  which  we  here  propose. 

We  are  certain  that  those  of  our  colleagues  who  are  also  interested 
in  the  systematics  of  the  Geophila  will  offer  corrections  on  the  basis  of 
new  facts:   nevertheless,  the  author  is  convinced  of  the  validity  of  the 
basic  classification  here  offered. 


It  is  commonly  admitted  that  conchological  featiires  in  general 
possess  only  little  taxonomic  value  and  that  the  area  of  their  applica- 
tion is  limited  to  the  familial  level,  and  more  often  to  the  generic 
or  specific  levels.   Thotigh  partly  true,  there  are  some  shell  features 
which  possess  a  considerahly  greater  informative  value  which  permits 
their  use  at  higher  taxonomic  ranks.   The  general  appearance  of  the 
shell,  as  well  as  the  presence  of  longitudinal  parietal  and  columellar 
plates,  are  such  characteristics.   Naturally,  when  we  speak  of  the  general 
shell  shape,  we  do  not  include  the  numerous  secondary  modifications . 

The  first  problem  to  present  itself  in  our  investigation  concerned 
the  nat;ire  of  the  original  Geophila  shell.   If  we  assume  that  the  order 
derived  from  ellobiid-like  ancestors — ^both  fossil  and  Recent  Ellobiidae 
are  characterized  more  or  less  by  a  monotypic  shell  of  elongated  form 
with  columellar  and  parietal  plates — then  we  must  accept  such  a  shell 
type  as  the  original  shape.   This  almost  \iniversally  accepted  opinion 
is  reinforced  by  the  fact  that  the  ancient  (and  although,  in  our  opinion, 
only  distantly  related)  Partulidae  and  Achatinellidae  do  indeed  possess 
precisely  such  a  kind  of  shell;  in  fact,  that  shape  is  characteristic 
for  the  lower  Orthurethra  as  well  as  for  the  lower  non-orthurethrous 
Geophila.   At  first  glance,  serious  deviations  from  this  shell  type  among 
the  lower  Geophila  are  seen  in  Pupilloidea  and  Sagdoidea:   i.e.  the 
Pyramidulidae ,  Valloniidae,  Strobilopsidae,  Sagdidae,  and  Thysanophoridae. 
However,  the  microhelicoid  shells,  characteristic  for  the  families  cited, 
are  "inscribed"  in  the  beginning  as  bulimuloid  shells,  which,  in  combina- 
tion with  anatomical  f ^atiires  ,  give  testimony  to  the  origin  of  those 
families  by  means  of  the  early  sexual  maturity  of  the  ancestral  form. 

and  consequently  in  the  early  interruption  of  shell  growth.   The  appearance 
protandry  (e.g.  in  the  Valloniidae) ,  which  leads  to  the  swift  reduction  of  the 
male  genital  system  after  having  performed  its  function,  is  another  consequence 
of  the  change  of  the  bulimuloid  shell  to  the  microhelicoid. 

In  the  almost  limitless  variations  of  the  shell  of  the  Geophila,  one  can 
detect  several  evolutionary  lines.   To  complete  the  survey  of  the  lower  mem- 
bers of  the  order,  we  note  a  tendency  toward  reduction  of  the  parameters  and 
retention  of  the  same  proportions  (e.g.  Pupillidae,  Cionellidae,  Vertiginidae , 
and  Orculidae).   This  tendency  is  bound  up  with  life  in  microhabits  ,  often — 
but  by  no  means  always — in  climatic  conditions  of  sharp  daily  temperature 
changes . 

The  other  tendencies  fit  in  with  the  natural  groupings,  that  is,  with 
the  suborders  proposed  in  the  present  work.   Thus  high,  many-whorled  shells, 
characteristic  of  most  of  the  Achatinidae  may  reach  a  very  large  size  in  a 
moist  tropical  climate.   The  inhabitants  of  forest,  and  secondarily  of  moun- 
tains, often  crawl  on  vertical  surfaces  and  frequently  with  the  ventral  side 
upward  and  evolved  a  long  narrow  shell  shape  ( Clausilioidea) .   The  columellar 
plates  in  such  cases  may  grow  stronger  since  they  are  the  point  of  support  for 
the  columellar  muscle  and  they  facilitate  the  evolutionary  direction  of  shell 
shape  (Likharev,  1962). 

In  the  lower  Oleacinina,  the  shell  initially  is  of  the  same  type,  but 
members  of  this  suborder  inhabit  a  large  number  of  biotypes  and  this  explains 
the  heterogeneity  of  their  shells.   The  greatest  degree  of  this  is  seen  in  the 
Streptaxidae,  among  which  can  be  fo\ind  almost  all  types  of  shell  shape  seen 
in  the  Geophila   (Zilch,  I961),  and  above  all,  in  the  unique  shell  type  of 
Gonaxis  with  its  displaced  or  distorted  axis.   The  Oleacinidae  possess  several 
unique  shell  features ,  rarely  encountered  elsewhere  than  in  the  suborder 

Oleacinina:  the  aperture  is  high  and  narrowed  in  its  upper  angular  part  and 
is  not  expanded  laterally:   the  combination  of  such  an  aperture  with  strongly 
flattened  whorls  and  weak  sutures  gives  the  shell  a  bullet-like  shape,  thus 
enabling  the  animals  to  crawl  in  the  soil  in  their  hunt  for  oligochetes,  their 
basic  food.   The  angular  area  of  the  apertural  region  is  an  adaptive  feature: 
in  this  region  the  pneumostome  is  displaced  and  the  mollusk  can  breathe  freely 
while  the  anterior  part  of  the  body  is  buried  under  the  solid  surface.   Charac- 
teristically, a  similar  shell  is  encountered  in  those  members  of  other  sub- 
orders which  are  also  closely  boimd  up  with  the  soil.   It  is  enough  to  observe 
that  a  predatory  life  style  in  the  Oleacinina  brings  about  a  tendency  to  shell 

reduction  as  in  the  case  of  two  other  higher  suborders,  the  Helixina  and  the 

,.    .     (1) 

Shell  reduction  itself  proceeds  along  two  evolutionary  pathways  associated 
with  either  l)  the  widening  of  the  last  whorl  and  consequently  of  the  aperture, 
or  2)  the  increasing  of  the  relative  height  of  the  aperture.   The  reduction  of 
the  shell  may  proceed  either  through  a  regularly  whorled  spiral  of  the  shell 
or  a  reduction  of  the  spire.   In  the  Helixina  the  attainment  of  the  slug 
shape  proceeds  along  the  first  path:   this  process  can  in  detail  be  followed 
in  the  Vitrinoidea.   Among  the  Oleacinina,  the  Testacellidae ,  however,  lose 
their  shells  by  passing  through  a  "succinoid"  stage.   The  increase  of  the  re- 
lative apertioral  height  can  be  traced  in  some  New  World  Oleacinidae:   Euglandina 
flammulata  H.B.B. — Pittieria  arborea  H.B.B. — Euglandina  pupa  H.B.B. — E.  deli- 
catula  (Shuttl.)  — E.  saxatilis  H.B.B. — Oleacina  camerata  H.B.B.  (Baker, 
19^1)  —  Strebelia  berendti  (Pfr.).  In  the  first  few  of  these  species  the 
aperture  height  is  35  0/0  of  the  shell  height,  in  the  last  it  is  93  0/0;  that 
is,  the  aperture  is  barely  shorter  than  the  shell. 

To  distinguish  them  from  the  generic  names  Helicina  and  Limacina. 

Similarly  in  the  lower  suborders  (Achatinina,  Oleacinina,  and  Pupillina) 
together  with  a  high  shell,  there  is  also  the  presence  of  coliomellar  plates 
which  may  either  disappear  or  grow  stronger  and  more  involved.   The  truncated 
columella  in  the  Oleacinidae,  Cionellidae,  and  many  Achat inellidae,  etc.  comes 
about  as  the  result  of  the  reduction  of  the  lower  part  of  the  columella  below 
the  plates:   the  lower  columellar  margin  in  this  case  corresponds  precisely 
to  the  columellar  plates . 

The  apertural  plates  mentioned  above  consist  of  contrasting  basal  and  pala- 
tal elements.   The  fact  is  that  these  two  groups  of  apertiiral  formations  have 
a  different  historical  natxire,  different  taxonomic  importance,  and  different 

As  far  as  the  basal  and  palatal  folds  are  concerned,  they  repeatedly  and 
independently  originated  in  all  the  suborders  which  are  themselves  characterized 
by  a  large  degree  of  diversity  and  variability  even  within  individual  species. 
Their  role  in  the  final  analysis  is  related  to  the  narrowing  of  the  opening  of 
the  aperture.   Their  beginning  is  bound  up  with  the  narrowing  of  the  last  whorl 
before  they  reach  their  definitive  state.   As  a  result  of  this  narrowing  the 
secretionary  surface  of  the  mantle  is  reduced  by  folding,  and  the  formation 
of  the  folds  leads  to  the  origin  of  the  apertural  armature. 

A  greatly  complicated  matter  is  seen  in  the  colijmellar  and  parietal  ele- 
ments.  We  emphasize  the  fact  that  there  is  no  important  distinction  between 
them:   in  many  ellobiids  the  boundary  between  the  columellar  and  parietal  aper- 
tural margins  is  indistinct  and  there  are  not  one  but  many  columellar  plate- 
lets (appearing,  apparently,  as  a  result  of  the  fracturing  of  one  of  them)  by 
which  the  upper  columellar  plates  actually  rest  on  the  parietal  wall.   In  con- 
trast to  the  basal-palatal  formations,  originating  either  at  the  very  end  of 
postembryogenesis ,  or  several  times  in  the  course  of  the  life  of  the  mollusk 

(Schileyko,  I96T)  ,  the  columellar  parietal  plates  appear  even  in  embryo- 
genesis  and  continue  to  lengthen  as  the  shell  grows.   Apropos  the  origin  of 
the  plates,  it  may  be  thought  that  the  single  origin  of  the  plates  is  a  con- 
sequence of  the  heterogeneity  of  the  ellobiid  ancestral  shell.   Such  hetero- 
geneity is  actually  present  in  some  ellobiids.   It  is  understood  that  because 
of  the  "break"  in  the  columella  at  the  jimction  of  the  embryonic  and  defini- 
tive whorls,  an  oblique  fold  appears,  which  extends,  as  the  shell  is  growing, 
to  the  aperture. 

The  special  function  of  the  col\jmellar  plates  apparently  consists  in  serving 
as  a  support  for  the  columellar  muscle.   In  cross  section  it  is  seen  that  this 
muscle  rests  on  the  upper  surface  of  the  plate:   if  there  are  2  plates,  the 
muscle  lies  between  them;  when  there  are  more  than  2  the  muscle  breaks  up  early 
into  branches,  each  of  which  occupies  a  niche  between  2  neighboring  folds.   The 
latter  condition  can  be  seen,  for  instance,  in  the  Urocoptidae  which  have  re- 
latively small  bodies  and  thus  the  problem  of  the  narrow  shell  is  especially 
severe.   Under  such  conditions  the  columellar  muscle  allows  the  shell  to  con- 
tinue to  grow  in  the  set  direction. 

The  principle  role  of  the  pariental  plate  is  to  force  itself  into  the 
mantle  cavity  and  incompletely  divide  it  into  right  and  left  halves.   We  re- 
call that  the  vascularization  of  the  right  and  left  halves  of  the  lung  is 
different:   the  respiratory  inflow  first  enters  the  left  half  of  the  cavity 
and  leaves  it  through  the  right:   thus  the  character  of  the  gas  exchange 
along  the  entire  plane  is  not  equal.   Apparently  the  more  primitive  division 
of  the  pulmonary  cavity  was  brought  about  to  regulate  the  air  current  and  to 
increase  the  effectiveness  of  breathing  —  a  creation  of  the  incompletely 
formed  partitions.   Later  such  a  method  of  dividing  the  lung  gave  way  to  a 
more  perfect,  active,  and  sensitive  method  with  the  help  of  the  mantle  folds 

in  aquatic  Basommatophora  (Sullivan  &  Cheng,  197^),  the  Helixina,  the 
Limaxina  and  thus,  independently,  in  the  remaining  higher  suborders  of 
the  Geophila. 

In  contrast  to  the  Achat inina,  Oleacinina,  and  Pupillina,  the  shell 
of  the  Helixina  and  Limaxina  was  originally  depressed  and  had  a  wide  um- 
bilicus as  often  found  in  the  Geophila.   However,  a  wide  unbilicus  is 
not  a  peculiar  phenomenon  among  the  Geophila;  it  usually  is  not  seen  in 
mollusks  with  a  shell  diameter  of  more  than  1  cm,  and  animals  with  such 
a  shell  live  in  strongly  shaded  areas.   In  similarly  shaped  shells  the 
relative  area  of  the  surface  is  large  and  its  usable  volume  is  small 
with  a  large  umbilicus  (Fig.  LA.)  .   While  as  the  umbilicus  narrowed,  the 
volume  which  was  taken  up  by  the  columellar  cavity  was  "given  over"  to 
the  accommodation  of  the  internal  organs  (Fig.  IB).   The  usual  mode  for 
the  narrowing  of  the  umbilicus  was  through  the  widening  of  the  whorls ; 
at  times  only  the  body  whorl  grew  wider  and  thus  the  interior  of  the 
shell  remained  a  cavity  in  commimication  with  the  interior  narrow  umbilical 

The  columellar  parietal  plates  in  some  African  and  Pacific  Endo- 
dontidae  are  not  homologous  to  those  of  the  Achatinina,  Oleacinina,  and 
Pupillina  which  are  formed  at  the  end  of  post-embryogenesis ;  they  may 
originate  conver gently. 

Many  Helixina  possess  thick-walled,  trochid-shaped  or  spherical  shells 
of  a  helicoid  appearance  without  any  apertural  armature;  however,  in  such 
cases,  when  the  apertxiral  teeth  are  present,  the  area  in  which  they  appear 
is  limited  to  the  aperture  and  the  adjacent  section  of  the  parietal  wall: 
they  do  not  run  along  the  columella  or  the  parietal  walls  to  any  distance 

inside  the  shell.  Further  in  this  suborder  there  exists  a  tendency  to  a 
thinning  of  the  shell,  and  the  repeated  appearance,  within  the  suborder, 
of  the  slug  form. 



We  have  already  discussed  the  data  and  elatorated  on  the  taxonomic 
and  evolutionary  significance  of  the  renopericardial  complex  (Schileyko, 
1976  a;  see  also  Minichev  &  Starohogatov,  1971).   We  will  therefore  here 
discuss  only  the  basic  proposals. 

In  embryogenesis  at  a  stage  comparable  to  the  veliger,  sigmurethry 
is  a  feature  even  in  orthurethral  forms  with  both  having  the  urethras  as 
open  furrows.   Basically  in  the  Orthurethra  the  kidney  differs  from  the 
kidney  of  related  Geophila  only  because  of  its  greater  length.   In  the 
more  progressive  groups  the  processes  of  the  shortening  of  the  kidney  and 
the  closing  of  the  urethra  take  place  as  an  evolutionary  parallelism  that 
is  not  necessarily  synchronous.   The  suborder  Pupillina  as  here  proposed, 
differs  from  the  Orthurethra  only  in  excluding  the  Partulidae  and  includ- 
ing the  Sagdoidea.   In  other  words,  the  Orthurethra  is  basically  a  natural 
taxon  but  contrasting  it  with  other  entities  at  the  same  rank  on  the 
basis  of  the  excretory  apparatus  is  not  justified. 

The  Heterurethra  evolved  as  a  result  of  the  decrease  of  shell  size 
and  the  consequent  shortening  and  widening  of  the  lung.   Here  the  kidney 
tiirns  either  clockwise  or  counter-clockwise  and  squeezes  itself  into  the 
posterior  wall  of  the  mantle  cavity,  finally  becoming  buried  inside  the 
visceral  sac. 

In  these  ways  the  featixres  of  the  renopericardial  complex  offer 
good  data  for  reconstructing  the  phylogenetic  relationships  in  super- 
families,  families,  and  sometimes  genera,  but  they  are  not  suitable  as 
basic  characteristics  for  suborders. 



The  positions  of  the  longitudinal  grooves  in  the  foot  are  used  for 
the  classification  of  only  the  higher  Geophila  —  the  Sigmurethra.   If 
one  takes  the  order  as  a  whole,  then  in  the  Achatinina,  Oleacinina,  and 
the  majority  of  Pupillina  the  foot  is  of  the  holopod  type  and  only  in 
some  Helixina  and  Limaxina  is  it  aulacopod. 

In  the  present  article  we  designate  the  furrows  rimning  along  the 
sole  as  the  pedal  furrows,  those  r-unning  along  the  edge  of  the  sole  as 
peripedal  furrows,  and  those  on  the  lateral  surface  as  suprapedal  furrows. 

What  is  the  function  of  the  furrows?  The  pedal  furrows,  seemingly 
facilitate  the  intensification  of  movement  by  strengthening  the  functional 
loading  of  weight  on  the  centrail  field  of  the  foot;  waves  of  muscular  con- 
traction in  this  case  move  along  the  central  field.   The  peripedal  and  supra- 
pedal  furrows  are  channels  along  which,  with  the  aid  of  a  ciliary  epi- 
thelium, mucus  is  driven  and  thus  they  perform  varied  protective  functions. 
It  is  natural  that  such  furrows  are  formed  as  a  result  of  life  in  arid  con- 
ditions, and,  in  the  ancestral  aquatic  shape,  permanent  furrows  were  absent 
in  the  foot .   The  first  furrows  which  appeared  in  the  dry-land  forms  were 
small  longitudinal  riffles,  issuing  from  below  the  buccal  flap  and  reach- 
ing the  anterior  half  of  the  body  (e.g.  Partulidae,  Fig.  2,  I,  II).   Later 
one  of  these  is  fractured  into  a  ring-shaped  peripedal  furrow,  occupying 
an  extremely  lateral  position  (e.g.  Ceriidae,  Subulinidae,  Clausiliidae , 
Pupillina  (Partly),  Helixina  (Fig.  2,  III)  or  a  suprapedal  furrow  (Vitrea, 
Fig.  2,  VIIl).  One  at  times  may  think  that  the  suprapedal  furrow  is  dis- 
placed and  thus  moves  to  an  extreme  marginal  position  (Fig.  2,  from  VIII 
to  III) . 


The  pedal  furrows  are  usually  homologous  (in  their  derivation)  to 
the  primary  suprapedal  or  peripedal  furrows.   The  only  exceptions  are 
those  very  rare  cases  when  a  single  medial  pedal  furrow  appears  ( Zoni- 
toides ,  Rhytida,  Fig.  2,  VI,  VII)  which  originates  in  a  fold  in  the  foot 
caused  when  the  animal  draws  itself  into  its  shell  and  thus  it  becomes 
isolated  (Schileyko,  19T2a) . 

Historically  the  succeeding  furrows  often  appear  later  than  the 
first  pair  of  peripedal  furrows,  and,  as  the  first  pair  "slips  down"  to 
the  sole,  they  occupy  their  place  (Fig.  2,  from  III  to  IV  to  V  and  from 
VIII  to  IX  to  IV  to  V). 

At  the  same  time  a  second  way  leading  to  the  origin  of  the  tripartite 
division  of  the  sole  is  conceivable:   namely  as  a  consequence  of  the  in- 
crease of  weight  on  the  central  part  of  the  holopod  sole.   Thus,  in  crawling. 
Achat ina  the  waves  of  contraction  do  not  take  in  the  whole  surface  of  the 
sole  (Fig.  2,  X) :   the  pedal  branches  of  the  columellar  muscle  are  strengthened 
in  the  central  field,  and  the  lateral  fields  are  moved  in  a  vertical  direction. 
In  other  words,  a  preadaptation  to  the  formation  of  a  tripartite  sole 
(Fig. 2,  from  X  to  XI)  is  apparent  here. 

Thus  (because  of  convergence)  the  pedal  incisions  are  not  suitable 
for  taxonomy  on  the  level  where  they  are  most  used.   Even  Pilsbry  admitted 
this  when  he  recalled  instances  where  the  members  of  the  Aulacopoda  had 
a  holopod  foot,  and  members  of  the  Holopoda  had  an  aulacopod  foot  (19^6: 

One  conclusion  which  could  be  drawn  from  the  data  is  extremely  im- 
portant for  a  further  discussion,  namely:   the  holopod  type  of  foot  in  any 
case  will  be  the  point  of  departure  leading  to  the  aulacopod.   The  latter 


arises  in  those  groups  whose  members  are  in  need  of  rapid  mobility :   either 
predators  or  slugs  upon  whose  speed  of  movement  depends  the  success  in  the 
passive  struggle  against  dessication. 



The  question  regarding  the  appearance  of  the  sexual  apparatus  in  the 
Geophila  is  of  prime  importance.   As  long  as  we  assume  that  the  Orthurethra 
are  the  most  primitive  group,  we  must  also  assume  the  same  for  their  sexual 
systems.   But  the  sexual  apparatus  of  the  orthiorethral  Pupillina  is  speciali- 
zed rather  than  archaic  and  cannot  be  regarded  as  the  point  of  departure  for 
the  other  types . 

If  we  leave  aside  the  numerous  cases  of  secondary  simplification,  then 
the  primary  simple  structure  of  the  genitalia  is  foimd  mostly  in  the  primi- 
tive Achatinina  and  Helixina:   in  the  first  case  the  Partulidae  and  Stropho- 
cheilidae,  in  the  second  the  Punctoidea  (in  the  Pupillina  and  Oleacinina, 
the  most  primitive  forms  do  indeed  possess  appendages  in  the  male  portion). 
Thus,  the  original  type  of  the  sexual  system  is  extremely  simple:   from  the 
alhumen  gland  issues  the  spermoviduct  (hermaphroditic  duct)  which  then  divides 
into  two  parts,  of  which  one  is  the  seminal  conductor  (vas  deferens)  which 
is  longer  than  the  vagina  and  forms  a  lateral  loop.   The  spermatheca  is 
still  not  differentiated  into  a  channel  ( spermathecal  duct)  and  reservoir 
( spermathecal  sac)  and  is  represented  only  by  an  elongated  sac  off  the  vagina. 

The  longest  appendage,  the  flagellum,  is  formed  in  a  basic  way  (Fig.  3). 
The  first  way  is  by  moving  the  point  where  it  enters  into  the  seminal  con- 
ductor (vas  deferens)  below  the  epiphallus.   The  second  mode  comes  about 
by  close  contact  of  the  distal  part  of  the  seminal  conductor  (vas  deferens) 
to  the  penis  and  the  subsequent  disappearance  of  the  partition  between  these 
channels.   In  general,  it  may  be  said  that  for  the  Pupillina  and  most  Helixina 
the  first  way  is  characteristic  while  the  second  is  for  the  Achatinina  and 



In  general,  in  the  Achat inina,  Endodontinia,  and  Limaxina  the  sexual 
apparatus  is  close  to  the  original  type.   But  in  the  superfamily  Achatinoidea 
there  is  one  peculiarity  rarely  seen  in  this  suborder:   the  seminal  conductor 
(vas  deferens)  is  in  close  contact  with  the  penis  and  epiphallus  and  often  is 
joined  to  these  channels,  covered  by  a  muscular  connective  tissue.   In 
r.ost  Achatinina  the  cover  is  clearly  seen  in  the  lower  part  of  the  penis, 
and  in  this  case  the  seminal  conductor  (vas  deferens)  pierces  the  said  cover 
or  passes  below  it  (Araujo,  1973;  Breure,  197^;  Mead,  1950).   Often  — 
and  in  various  Helixina  —  a  penial  sheath  is  formed,  but  here  the  seminal 
conductor  (vas  deferens)  either  remains  free  or  is  drawn  to  the  atrial  area 
by  sheaves  and  does  not  penetrate  its  own  cover . 

If  we  now  turn  to  the  Pupillina  we  see  the  unique  genitalia  of  this  sub- 
order sharply  delineated.   The  penis  has  two  appendages  (Fig.  k) :      a  small 
caecum  and  an  appendix  consisting  of  5  sections.   In  a  full  view  we  distinguish: 
1.   a  basal  cylindrical,  swollen  section  (Al);   2.   a  spherical  section 
separated  from  the  base  by  a  sphincter  (A2);   3.   a  short,  narrow  cylindri- 
cal structure  (A3);   ^.   a  thin  tube  with  muscular  walls  (AU);   5.   a  widened, 
lengthened  ampule  (A5)  (Schileyko,  1976b).   Based  on  this  —  and  many  authors, 
without  providing  data,  homologize  the  penial  appendix  with  the  stylophora 
Cdart  sac)  and  mucus  glands  of  the  higher  Geophila  (Albers,  i860;   Forcart, 
19^0;  Ihering  l892;  1909;  Wiegmann,  1900)  —  we  must  note  that  the  appendage 
fulfills  a  stimulating  function  before  coitus.  There  is  also  the  supposition 
that  the  appendage  is  involved  in  the  reception  of  the  partner's  spermato- 
phore.   As  to  the  caecum,  Forcart  (19^0)  ascribed  to  it  the  function  of  the 
mechanical  place  of  the  retention  of  the  spermatophore  before  copulation. 


The  fmiction  of  the  epiphallus  (and  flagellum)  is  the  same  as  in  other 
groups  with  spermatophores ;   namely,  the  formation  of  a  cover  for  the 

It's  easy  to  understand  the  role  of  the  caecum  when  an  individual  with 
the  spermatophore  still  contained  inside  the  penis  is  opened:   the  cover  of 
the  latter  has  a  "spur"  formed  in  the  cavity  of  the  caecum  corresponding  to 
the  shape  of  the  cavity.   The  spur  apparently  is,. an.  anchor  wbi.Gh_.preyeOts 
the  spermatophore  from  leaving  the  sexual  system  earlier  than  is  necessary. 
The  spermatophore  itself  has  a  gutter  or  furrow  "over"  which  is  drawn  a 
thin  film,  but  the  film  is  absent  in  the  posterior  part  (Fig.  5).   In  a 
histological  examination  of  individuals  of  some  Buliminidae  species, 
collected  immediately  before  copulation,  sperm  was  detected  inside  A5 . 
It  is  apparent  that  this  could  not  be  allosperm  (sperm  of  the  partner); 
firstly,  because  coitus  had  not  yet  taken  place,  and  secondly,  because  there 
is  a  spermatheca  for  the  reception  of  the  allosperm.   Consequently  the 
sperm  noted  was  autosperm  and  thus  A5  serves  as  a  supplementary  depot  for 
the  autosperm  where  the  latter  enters  before  coitus  and  possibly  there  also 
awaits  the  period  of  ripening. 

To  sum  up,  in  an  animal  ready  for  coitus  the  sperm  is  collected  inside 
A5,  and  the  spermatophore  is  formed  already  but  it  is  empty.   In  copulation 
the  penis  is  twisted,  Al  and  A2.   The  spermatophore  is  delivered  to  the  partner, 
and  when  the  larger  part  of  it  has  already  left  the  penis,  the  sperm  is  forced 
into  the  cavity  of  the  spermatophore  cover  by  contraction  of  the  musculature 
in  A3  and  aU .   A2  serves  as  the  regulator  of  the  entrance  of  sperm  into  the 
spermatophore  since  it  is  provided  with  one  or  two  sphincters  and  papillae. 

This  is  the  situation  in  many  Palearctic  Pupillina.   Extraordinarily 


interesting  modifications  of  this  type  of  genitalia  appear  in  the  Pacific 
Achat inellidae  (Cooke  &  Kondo,  I960):  in  members  of  some  groups  there  is 
a  substitution  of  the  penial  appendix,  as  a  result  of  which  the  latter  is 

To  account  for  this  modification  of  the  male  apparatus  among  Pacific 
taxa,  one  expects  to  see  some  reduction  of  the  penial  appendix  among  Paleartic 
families.   Indeed,  there  is  only  a  single  family  where  this  occurs:   the 
Orculidae  (Schileyko,  1976b;  Steenberg,  1925)  —  which  in  any  case  shows 
a  large  degree  of  uniqueness  in  the  organization  of  the  male  sexual  structures. 

Furthermore  in  the  Geophila  the  presence  of  a  more  or  less  complicated 
structure  of  the  quadrivia  is  characteristic,  in  the  composition  of  which 
there  is  a  fertilization  chamber,  a  spermatheca  and  also  incurrent  and  ex- 
current  channels   (L.  Schileyko  and  A.  Schileyko.  1975;  Flasar ,  1967;  Lind, 
1973;  Van  Mol,  1971).   As  far  as  is  known,  the  only  group  where  the  quadrivia 
are  represented  simply  by  a  bend  of  the  hemaphroditic  channel  is  the  Partu- 
lidae,  and  this  bears  witness  to  the  primitiveness  of  this  group  when  compared 
to  the  other  Geophila. 

Once  again  we  emphasize  the  fact  that  the  simplest  sexual  apparatus 
as  a  whole  is  found  in  Achatinina  (Fig.  6)  and  thus  this  simplicity  is  of 
a  primary  character.   The  most  complicated  element  —  the  penial  papillae  — 
is  only  seen  from  time  to  time  in  this  group  (Fig.  6,  V)  independently  of  the 
formation  of  analogous  structures  in  other  suborders . 

The  sexual  apparatus  of  the  higher  Helixina  —  Helixinia  and  Zonitinia 
and  also  Limaxina  has  an  entirely  different  aspect.   In  these  taxa  the  primary 
presence  of  vaginal  appendages  is  characteristic  —  the  sarcobelum,  stylo- 
phores  or  other  appendages  like  the  atrial  gland  in  Milacidae.   The  most 


commonly  associated  organs  of  the  stylophores  are  the  mucous  glands  although 
at  times  the  sarcobelum  is  also  provided  with  glands. 

It  may  be  thought  that  all  these  appendages  came  about  as  a  result  of 
a  differentiation  of  non-specialized  glandular  tissue  in  the  lower  part  of 
the  vagina,  since  their  formation  took  place  no  fewer  than  3  times:  l) 
when  the  h  stylophores  and  mucous  glands  are  arranged  like  a  wreath  around 
the  vagina  (Humboldtianidae) ;  2)  when  the  h  stylophores  are  arranged  in  2 
rows  (Hygromioidea) i  in  both  cases  the  number  of  stylophores  is  reduced  to 
one;   3)  the  position  of  a  single  sarcobelum. 

As  for  the  male  portion  in  the  Helixinia,  in  the  Endodontinia  the 
penis  is  simple,  at  times  sac-like,  often  with  internal  appendages  (Climo, 
1969,  1970,  1971);  Riedel  and  Wiktor,  197^;  Solem,  1970).   The  only  external 
appendage  —  often  absent  —  is  the  flagellum.   The  penis  of  the  Helixinia, 
especially  in  the  Vitrinoidea,  is  more  complex.   Here,  in  addition  to  the 
flagellum,  there  is  also  a  caecum,  not  homologous  to  the  caecum  of  the  Pupil- 
lina.   In  answer  to  the  complicated  shape  of  the  spermatophore  cover,  the 
cavity  of  the  flagellum  is  also  complicated.   In  a  series  of  Recent  forms, 
one  can  trace  the  process  of  the  simplification  of  the  male  portion  through 
the  disappearance  of  the  caec\im;   the  flagellum  likewise  shortens  and  may 
be  reduced,  and  the  surface  of  the  spermatophore  is  often  also  simplified. 
In  Pseudoaustenia  ( Ariophantidae)  the  flagellum  also  disappears,  but  the  in- 
ternal surface  structures,  characteristic  of  the  species,  have  an  internal 
epiphallus  (Van  Mol,  1973).   Among  various  taxa  of  the  Helixinia,  as  also 
Eimong  the  Achatinina,  the  formation  may  be  seen  of  a  penial  sheath  (many 
Euconulidae,  Helicoidea,  Vitrinoidea,  etc.). 

Finally  there  is  one  other  important  element  in  the  male  portion  —  the 


penial  papillae.   Very  often  the  papillae  are  present  in  the  Helixinia,  and 
in  some  cases  even  2  papillae  are  formed  (Schileyko,  1972"b),  but  these  may 
appear  independently  in  members  of  all  the  other  taxa. 

In  conclusion  we  note  that  there  are  no  characteristics  in  the  sexual 
apparatus  sufficient  to  clearly  differentiate  3  infraorders  within  the 
Helixinia,  even  if  we  do  not  consider  the  reduction  of  characteristics  which 
may  lead  in  time  to  a  formal  resemblance  based  on  negative  characteristics. 
Thus  several  Helicoidea  ( Oreohelicidae,  Bradybaenidae)  characteristically 
have  one  stylophore  on  which  the  mucous  glands  are  located;  however,  on  the 
external  sarcobellum  in  Elaphroconcha  (Ariophantidae) ,  k   mucous  glands  are 
similarly  located. 

We  now  proceed  to  the  diagnostic  characteristics  of  the  suborders  and 
the  infraorders,  and  we  will  also  attempt  to  define  their  limits  and  size. 


Order  Geophila  Ferussac,  l8l2 
Suborder  Achatinina  Schileyko,  new 

The  shell  is  achatinoid,  bulimuloid,  or  elongate-fusiform  in  shape  with 
columellar  lamellae.   Longitudinal  parietal  lamellae  are  absent  or  present 
only  in  the  last  whorl;   the  aperture  often  lacks  teeth.   The  sexual  system 
is  without  appendages  except  for  a  flagellum  on  the  penis;  often  there  is  a 
close  connection  of  seminal  duct  (vas  deferens)  to  the  penis  and  a  penial 
sheath  appears.   The  kidney  is  shortened  with  the  urethras  closed  to  differ- 
ing degrees.   The  foot  is  holopod.   Obligatory  predators  are  not  found  in 
the  suborder.   Oviparous,  very  rarely  is  ovoviviparity  observed. 

1.  Superfamily  Achatinoidea 

(Megalobulimidae  Leme,  1973,  Strophocheilidae  Thiele,  1926; 
Odontostomidae  Pilsbry  et  Vanatta,  I898;  Bulimulidae  Tryon, 
1867;  Amphibulimidae  Crosse  et  Fischer,  l873;  Anadromidae 
Zilch,  1959;  Achatinidae  Swainson,  l8i+0;  Dorcasiidae  Conolly, 
1915;  Acavidae  Pilsbry,  l895 ;  Clavatoridae  Thiele,  1926). 

2.  Superfamily  Subulinoidea 

(Subulinidae,  Crosse  et  Fischer,  l877;  Ferussaciidae 
Bourguignat ,  l883;  Stenogyridae  Wenz  ,  1923). 

3.  Superfamily  Clausilioidea 

(Megaspiridae  Pilsbry,  I90U;  Urocoptidae  Pilsbry  et  Vanatta, 
1898;  Clausiliidae  Morch,  lQ6h;    Filholiidae  Wenz,  1923). 
h.      Superfamily  Par tulo idea 

(Partiaidae  Pilsbry,  1900 ). 


Suborder  Oleacinina  Schileyko,  new 
For  the  most  part  the  shell  is  elongate  with  columellar  lamellae. 
Longitudinal  parietal  lamellae  are  wanting  though  the  aperture  at  times  has 
teeth.   The  female  genitalia  do  not  have  appendages  and  in  the  male  portion, 
the  penis  is  simple  or  with  1-2  appendages.   The  foot  is  holopod.   These 
obligatory  predators  are  oviparous. 

1.  Superfamily  Testacelloidea 

(Spiraxidae  Baker,  1955;  Oleacinidae  Adams,  1855; 
Testacellidae  Gray,  18A0) 

2.  Superfamily  Streptaxoidea 

(Streptaxidae  Gray,  1860) 

Suborder  Pupillina  Schileyko,  new 
The  shell  is  bulimuloid,  rarely  depressed  or  microhelicoid  with  colu- 
mellar lamellae.   The  parietal  lamellae  are  characteristic  and  the  aperture 
often  has  parietal  teeth.   Female  genitalia  are  without  appendages  and  the 
penis  primarily  has  a  caecum  and  appendix  consisting  of  5  sections.   The  foot 
is  holopod.   With  the  exception  of  the  Sagdoidea,  there  are  no  obligatory 
predators  in  the  suborder.   Oviparous,  very  rarely  ovoviviparous . 

1.  Superfamily  Cerioidea 

(Ceriidae  Fleming,  1818) 

2.  Superfamily  Achatinelloidea 

(Dendropupidae  Wenz,  1938;  Achatinellidae  Gulick,  1873; 
Orculidae  Pilsbry,  1918) 

3.  Superfamily  Cionelloidea 

(Amastridae  Pilsbry,  1911,  Cionellidae  Clessin,  1879) 


4.  Superfamily  Pupil loidea 

(Pupillidae  Turton,  1831;  Buliminidae  Clessin,  1879; 
Chondrinidae  Steenberg,  1925;  Pyramidulidae  Wenz,  1923; 
Valloniidae  Morse,  1864;  Vertiginidae  Fitzinger,  1833; 
Strobilopsidae  Pilsbry,  1918) 

5.  Superfamily  Sagdoidea 

(Thysanophoridae  Pilsbry,  1926;  Sagdidae  Pilsbry,  1895) 

Suborder  Helixina  Schileyko,  new 
The  shell  is  flat  to  turbinate,  often  thin  and  translucent  and  without 
columellar  lamellae.   There  are  no  longitudinal  parietal  lamellae  and  the 
apertural  margin  is  simple  or  with  teeth.   The  female  portion  of  the  geni- 
talia and/or  the  atrium  primarily  bears  variable  appendages  or  is  provided 
with  a  perivaginal  gland.  The  penis  has  a  flagellum;  there  are  often  other 
appendages  on  the  male  genitalia.   The  kidney  is  shortened,  the  primary 
urethra  is  closed,  the  secondary  urethra  is  either  completely  or  partially 
closed.   The  foot  is  holopod  or  aulacopod.   Together  with  herbivores 
there  are  some  obligatory  predators.   Oviparous  or  ovoviviparous. 

Infraorder  Endodontinia  Schileyko,  new 
The  shell  is  depressed  with  a  wide  umbilicus  and  often  with  strong 
radial  sculpture.   The  aperture  often  has  teeth.   The  genitalia  are 
simple  with  a  more  or  less  elongate  vagina,  at  times  with  a  glandular 
section.   The  penial  papillae  are  absent.   The  foot  is  holopod  or  aulaco- 


1.  Superfamily  Punctoidea 

(Endodontidae  Pilsbry,  1894;  Polygyridae  Pilsbry,  1895; 
Helicodiscidae  Pilsbry,  1927;  Punctidae  Morse,  1864; 
Pleurodiscidae  Wenz,  1923;  Systrophiidae  Thiele,  1926) 

2.  Superfamily  Thyrophorelloidea 

CThyrophorellidae  Girard,  1895) 

Infraorder  Helixinia  Schileyko,  new 
The  shell  is  flat  to  spherical  with  the  umbilicus  narrowed  to  varying 
degrees.   Radial  rib-like  sculpture  on  shell  is  exceptional  and  apertural 
teeth  are  rare.   The  genitalia  are  complicated  with  a  perivaginal  gland, 
sarcobellum,  stylophores,  or  mucous  glands  with  the  latter  usually  connected 
to  the  stylophores  or  the  sarcobellum.   Penial  papillae  are  generally 
present.   The  foot  is  holopod  or  aulacopod. 

1.  Superfamily  Gastrodontoidea 

(Euconulidae  H.B.  Baker,  1928;  Gastrodontidae  Tryon,  1866;) 

2.  Superfamily  Rhytidoidea 

(Rhytididae  Pilsbry, 1893;  Haplotrematidae  H.B.  Baker,  1925; 
Chlamydephoridae  Cockerell,  1935) 

3.  Superfamily  Vitrinoidea 

(Helicarionidae  Bourguignat,  1888;  Trochomorphidae 
Mollendorff,  1890;  Urocyclidae  Simroth  1889;  Ariophantidae 
Godwin-Austen,  1888;  Vitrinidae  Fitzinger,  1833) 

4.  Superfamily  Arionoidea 

(Otoconchidae  Cockerell,  1893;  Arionidae  Gray,  1840; 
Phylomycidae  Gray,  1847) 


5.  Superfamily  Sphincterochiloidea 

(Sphincterochilidae  Zilch,  1959) 

6.  Superfamily  Helicodontoidea 

(Helicodontidae  Hesse,  1918) 

7.  Superfamily  Helicoidea 

(Humboldtianidae  Pilsbry,  1939;  Helicidae  Rafinesque,  1815; 
Helminthoglyptidae  Pilsbry,  1939;  Bradybaenidae  Pilsbry  1939; 
Corillidae  Pilsbry,  1905;  Oreohelicidae  Pilsbry,  1939; 
Camaenidae  Pilsbry,  1895;  Ammonitellidae  Pilsbry,  1930) 

8.  Superfamily  Hygromioidea 

(Hygroraiidae  Tryon,  1866) 

Infraorder  Zonitinia  Schileyko,  new 
The  shell  is  depressed,  zonitoid,  with  the  umbilicus  narrowed  to 
varying  degrees;  a  clear  tendency  to  shell  reduction  and  slug  form  is 
present.   Sculpture  is  weak;  apertural  teeth  are  absent.   The  female  portion 
of  the  genitalia  has  a  perivaginal  gland  or  atrial  gland.   The  penial 
papillae  are  mostly  absent  or  closed.   The  foot  is  aulacopod. 

1.  Superfamily  Zonitoidea 

(Zonitidae  Mbrch,  1864;  Daudebardiidae  Pilsbry, 1908) 

2.  Superfamily  Parmacelloidea 

(Parmacellidae  Gray,  1860;  Milacidae  Germain,  1930) 

Suborder  Limaxina  Schileyko,  new 
The  shell  is  represented  by  a  plate  or  separate  calcareous  grains. 
The  female  portion  of  the  genitalia  is  without  appendages;  the  penis  is 


likewise  without  appendages  though  at  times  a  short  flagellum  is  present. 
The  internal  structure  of  the  penis  at  times  is  very  complicated.  The 
kidney  is  very  variable  and  bears  a  relationship  to  the  reduction  of  the 
shell;  the  urethras  are  closed.  The  foot  is  aulacopod.  These  oviparous 
animals  are  herbivores  or  predators. 

Infraorder  Trigonochlamydinia  Schileyko,  new 
The  mantle  is  small,  most  often  without  a  hood  and  sharply  posterior. 
The  penis^.is  tube-like, smooth  internally;  spermatophores  rest  on  glandular 
pillows  and  their  shapes  do  not  correspond  to  the  shape  of  the  penis.   Obli- 
gatory predators. 

1.   Superfamily  Trigonochlamydoidea 
(Trigonochlamydidae  Hesse, 1882) 

Infraorder  Limaxinia  Schileyko,  new 
The  mantle  is  well  developed,  with  a  hood  most  often  on  the  anterior 
part  of  the  body.  The  penis  is  elongate  or  sac-like  and  spermatophores 
are  not  present.   Herbivores. 

1.   Superfamily  Limacoidea 

(Boetgerillidae  Van  Goethem,  1972;  Limacidae  Rafinesque,  1815; 
Agriolimacidae  Wagner,  1935) 



From  the  data  available  it  appears  that  in  the  Geophila  there  are  two 
basic  phylogenetic  branches:  A)   shell  high,  with  columellar  (and  often  with 
parietal)  lamellae;  female  genitalia  without  appendages.   B)   shell  depressed, 
without  columellar  parietal  lamellae;  female  genitalia  show  an  inclination 
to  the  formation  of  various,  at  times  very  complicated,  appendages. 

The  first  group  includes  the  Achatinina,  Oleacinina,  and  Pupillina;  in 
the  second  are  the  Helixina  and  Limaxina.   The  apportionment,  however,  is  not 
into  2  but  rather  5  suborders  and  this  is  clearly  seen  from  the  early  diversi- 
fication of  the  corresponding  phylogenetic  branches  (Fig.  7). 

Near  the  base  of  the  first  phylogenetic  branch  stands  the  Partulidae. 
The  reasons  for  thus  placing  the  family  are  the  following:   the  shell  is 
primitive,  although  it  also  has  such  progressive  features  as  an  absence  of 
parietal  lamellae.   The  kidney  is  orthurethral,  but  one  can  already  note  the 
differentiation  into  sac-like  proximal  and  narrow  distal  divisions  (Fig.  8B) . 
The  foot  is  extraordinarily  archaic  (see  above).   The  sexual  apparatus  is 
also  extraordinarily  and  primitively  simple  (Fig.  8V);  the  simplicity  is  not 
a  result  of  the  reduction  of  any  appendages;  the  seminal  tube  (vas  deferens) 
is  not  connected  directly  with  the  penis  and  a  penial  sheath  is  lacking. 
One  may  think  that  the  progressive  (derived)  features  of  the  Partulidae  are 
bound  up  with  oviparity;   individuals  of  the  Partulidae  from  time  to  time 
lay  1-2  relatively  large  eggs,  the  diameter  of  which  we  compare  to  the  dia- 
meter of  the  aperture  of  the  mature  mollusk.   It  is  clear  that  when  laying 
eggs  of  such  a  size  --  large  enough  to  contain  the  developing  young  --  the 
formation  of  any  hard  structures  such  as  dentition  in  the  aperture  would  be 
hampering.   Kondo  fj  Burch  (1972)  also  emphasize  the  antiquity  of  the  group. 


These  circumstances  force  one  to  see  in  the  Partulidae  a  taxon  which 
is  close  to  the  basic  common  stem  from  which  the  Achatinina,  Oleacinina,  and 
Pupillina  were  derived.  The  closer  phyletic  association  of  the  Partulidae 
especially  to  the  Achatinina  is  based  primarily  on  the  sexual  apparatus. 
This  is  one  of  the  arguments  for  regarding  the  Achatinina  as  the  most  primi- 
tive suborder  and  not  the  Pupillina  as  is  now  believed. 

This  conclusion  is  confirmed  by  other  arguments.  Thus,  in  Achatinina, 
in  addition  to  Partulidae,  there  occurs  an  intensively  developing  process  in 
the  formation  of  sigmurethry,  beginning  for  the  most  part  with  the  shortening 
of  the  kidney.   Beginning  mesurethry  in  this  way  is  a  feature  of  the  lower 
Achatinina  (Megalobulimidae,  Strophocheilidae,  etc.).  There  is  also  another 
way:   (of  evolving  sigmurethry)  the  closing  of  the  urethra  without  noticeably 
shortening  the  kidney  (Antidrymaeus,  Fig.  6G).   Leme  (1973)  suggested  that  the 
Megalobulimidae,  proposed  by  him,  is  one  of  the  most  primitive  groups  of  the 
order,  and  this  once  again  emphasizes  the  correctness  of  our  view  regarding 
the  systematic  and  phylogenetic  placement  of  the  Achatinina.  The  Megalobuli- 
midae is  the  ancestor  of  the  ancient  form  from  which  the  Partulidae  also 
originated,  but  the  Megalobulimidae  is  more  progressive  or  advanced  in  the 
structure  of  the  kidney. 

One  can  not  doubt  the  unity  of  the  conchologically  strongly  characterized 
Clausilioidea,  if  one  omits  the  fossil  Filholiidae.   But  even  the  sinistral 
Filholiidae  already  bear,  in  a  weakly  developed  form,  the  features  specifi- 
cally peculiar  to  the  Clausiliidae:   an  apertural  fold  and  a  clearly  defined 
sinulus.  The  similarity  and  the  simple  arrangement  of  the  genitalia  of  the 
Clausiliidae  and  Urocoptidae  developed  convergently  and  independently  from  the 
Megaspiridae.   If  in  Clausiliidae  the  development  of  the  shell  was  by  way  of 


strengthening  the  apertural  armament,  then  in  Urocoptidae  it  took  place  in 
a  basically  different  manner:   through  the  differentiation  of  the  columellar 
lamellae.   Moreover,  the  Urocoptidae  is  more  progressive  in  the  organization 
of  the  excretory  apparatus. 

The  shell  of  the  Pupillina  is  more  advanced  in  general  features.  The 
following  should  be  added.   We  propose  the  following  points  which  diverge 
even  more  from  previous  schemes  and  generally  accepted  opinions:   1)   the 
union  in  one  superfamily  of  the  fossil  Dendropupidae,  the  eastern  Mediter- 
ranean Orculidae,  and  the  Hawaiian  Achatinellidae;   2)  the  introduction 
into  the  suborder  Pupillina  of  the  Ceriidae  and  the  superfamily  Sagdoidea. 

To  the  first  of  these  two  points  the  following  considerations  should 
be  stated.   Knight  (Moore,  1960)  places  Dendropupidae  in  the  Cyclophoridae 
together  with  2  other  genera.   However,  it  is  obvious  that  as  a  matter  of 
fact  Dendropupa  properly  belongs  with  the  Geophila  and  is  not  related  to 
Maturipupa  and  Anthracopupa.   Therefore,  when  we  speak  of  the  Dendropupidae 
we  have  only  the  real  Dendropupa  in  mind.   Thus,  the  families  Dendropupidae, 
Orculidae,  and  Achatinellidae  are  united  by  the  shape  of  the  shell  and, 
most  fundamentally,  by  the  presence  of  well  developed  parietal  lamellae 
running  from  the  first  whorl  to  the  last.   As  for  the  internal  features, 
the  Orculidae  and  the  Achatinellidae  are  the  only  Pupillina  in  which 
the  variations  of  the  copulatory  apparatus  can  not  easily  be  ascribed  simply 
to  a  reduction  of  the  penial  appendix  or  the  origin  of  small  partial  changes, 
but  they  bear  a  'creative'  character;  i.e.,  different  elements  are  developed; 
they  alter  the  surface  appearance  and  the  internal  structure,  and  by  no 
means  are  they  cast  in  the  static  mold  as  in  the  other  Pupillina  [cf.  Cooke 
5  Kondo,  1960;  Schileyko,  1976b).   Therefore,  although  one  can  not  speak  of 


an  immediate  closeness,  there  are  some  reasons  to  place  them  in  the  same 
superfamily,  Achatinelloidea. 

As  for  the  Ceriidae  (Fig.  9),  the  open  secondary  urethra  testifies 
to  the  definite  primitiveness  of  the  group;  the  case  of  the  sexual  appara- 
tus is  somewhat  more  complicated.   Firstly, the  presence  of  a  diverticulum 
in  the  sperraatheca  is  for  the  Pupil lina  much  more  characteristic  here  than 
for  the  other  groups.   Secondly,  the  support  or  attachment  of  the  penial 
retractor  muscle  to  the  flagellum  or  penial  appendix,  (Fig. 9,  PA),  is  a 
rare  phenomenon  and  occurs  only  in  aberrant  species  of  the  higher  Geophila. 
From  this  it  follows  that  the  long  appendage  to  which  the  retractor  in  Cerii- 
dae is  fastened  is  apparently  the  rudiment  or  derivative  of  the  penial  ap- 
pendix.  If  this  is  true,  then  the  family  must  be  placed  in  the  Pupillina. 
If  one  tries  to  find  a  place  for  the  Ceriidae  outside  the  Pupillina,  this 
must  be  only  with  the  Achatinina.  At  the  same  time  the  unique  Ceriidae 
must  be  separated  into  a  monotypic  superfamily,  the  Cerioidea. 

In  regard  to  the  Sagdoidea  we  must  say  that  there  are  two  serious 
objections  to  including  them  in  the  Pupillina:   the  depressed  shell  and  the 
sigmurethran  type  of  excretory  apparatus.  Actually  the  depressed  shell 
is  not  typical  in  the  Pupillina;  the  flat  shell  of  Hendersoniella  does  not 
prevent  the  association  of  that  genus  with  the  Urocoptidae  and  the  cylindri- 
cal Cylindrus  is  associated  with  the  Helicidae.   In  addition, there  exists 
the  possibility  of  associating  the  strongly  flattened  shell  of  the  type 
of  the  sagdid  Lacteoluna  with  the  high  shells  of  the  Pupillina  by  way  of 
the  high  cupola-shaped  shells  of  the  West  Indian  Sagda.  As  for  excretory 
apparatus,  one  may  determine  that  the  secondary  urethra  in  different  groups 
is  not  completely  closed,  and  the  length  of  the  kidney  in  the  Sagdoidea 


is  2-3  times  longer  than  the  pericardium  and  takes  in  almost  half  the  roof 
of  the  mantle  cavity;  in  other  words,  the  Sagdoidea  is  not  so  greatly  dif- 
ferent from  the  other  Pupillina  in  this  respect.   In  this  connection  it 
is  necessary  to  take  into  account  the  fact  that  all  Sagdoidea  are  predators 
and  such  a  life  style  always  strongly  stimulates  the  formation  of  sigmurethry. 

At  the  same  time  the  presence  of  a  penial  appendix  is  characteristic  of 
the  Pupillina,  and  moreover,  only  the  Pupillina.   Some  Sagdoidea  (e.g.  Lacteo- 
luna)  have  an  appendix  of  sufficiently  typical  structure  (Pilsbry,  1940)  . 

Let  us  pass  on  to  an  examination  of  the  suborder  Oleacinina.   This  taxon, 
overlooking  for  the  moment  the  great  conchological  variability  of  the  Strep- 
taxidae,  has  a  sufficiently  characteristic  shell  and  a  spiral  columellar 

Here  it  is  necessary  to  digress  and  once  again  strongly  emphasize  the 
fact  that  if  we  take  into  account  all  modifications  of  shell  shape  (in  this 
case  for  the  Streptaxidae) ,  it  would  be  impossible  to  establish  a  differential 
diagnosis  for  most  families  or  for  that  matter  the  majority  of  the  genera 
of  the  Geophila,  not  to  mention  taxa  of  higher  rank.  Therefore,  schematic 
generalization  is  always  unavoidable.   When  one  mentions  the  name  of  any 
taxon  there  appears  in  the  mind  of  the  reader  not  some  kind  of  definite  pic- 
ture, but  rather  a  sort  of  generalized  image  of  a  shell  shape  which  neverthe- 
less contains  most  of  the  characteristic  features  of  the  taxon  as  a  whole. 
Such  a  generalized  picture  nevertheless  presents  a  very  real  idea  of  the 
group.   We  had  such  considerations  in  mind  when  we  added  figures  of  the 
shells  in  Figure  7. 

Let  us  return  to  the  examination  of  the  Oleacinina.   This  suborder, 
in  spite  of  all  the  variations  of  the  shell,  is  quite  compact.   The  repro- 


ductive  systems  of  the  Spiraxidae  and  the  Streptaxidae  do  not  differ  greatly 
and  the  chitinous  hooks  which  occur  inside  the  penis  in  several  Streptaxidae 
are  doubtlessly  later  derived  acquisitions. 

The  generally  simple  structure  of  the  distal  portions  of  the  sexual 
system  is  characteristic  for  all  Oleacinina.   The  only  complication  in  the 
Oleacinidae,  for  instance,  is  the  presence  of  one  --  rarely  two  --  sac-like 
growths  on  the  penis  (Fig.  10);  in  some  of  the  Streptaxidae  a  penial  appendage 
may  also  be  present. 

Let  us  proceed  to  the  higher  suborders  --Helixina  and  Limaxina.   First 
of  all  we  must  point  to  the  widely  held  erroneous  opinion  that  the  Helicidae  -- 
more  definitely  the  genus  Helix  --represents  the  highest  development  of  the 
Geophila.   The  slugs  without  doubt  are  the  biologically  most  progressive 
shell-bearing  raollusks,  since  the  shell  itself  is  a  hindrance  to  biological 
advance,  because  any  external  stimulus  brings  forth  a  single  stereotyped 
reaction:   the  animal  pulls  back  into  the  shell.  The  mollusks  without  a  shell 
must  move  more  rapidly  and  possess  a  certain  level  of  neural  organization  in 
order  to  react  adequately  to  any  kind  of  stimulus.   Such  was  shown  by  Zs.-Nagy 
§  Sakharov  [1970)  who  found  a  much  stronger  development  of  the  web  of  synapses 
in  the  procerebrum  of  Limax  than  in  Helix.  This  is  further  exhibited  in  the 
fact  that  in  many  slugs  or  semi-slugs  there  is  a  tripartite  division  of  the 
sole  which  insures  a  greater  speed  of  movement.   Of  course,  this  does  not 
mean  that  such  a  sole  is  a  necessary  diagnostic  feature  of  this  group. 

In  Fig.  7,  the  infraorder  Endodontinia  is  treated  like  the  main  (stem) 
group  in  relationship  to  the  other  Helixina.  This  treatment  is  supported  by 
the  following  facts  and  considerations.  The  shell  of  the  Endodontidae  and 
related  groups  is  little  specialized  and  may  be  regarded  as  the  origin  of 


almost  all  the  conchological  types  of  the  higher  Geophila.  The  foot  is  holo- 
pod,  the  kidney  sigmurethran,  although  in  several  Helicoidea  the  secondary 
urethra  may  be  partially  or  completely  closed  (Schileyko,  1976a;  Wurtz,  1955) . 
The  sexual  apparatus  has  a  primary  simple  structure,  and  it  is  precisely  on 
account  of  that  low  level  of  specialization  that  the  ancient  Endodontinia 
proved  to  be  so  plastic  that  they  provided  the  origin  of  the  phylogenetic 
branch  of  the  three  infraorders  here  delineated.   The  basic  directions  of 
specialization  may  be  characterized  as  follows: 

1.  A  tendency  to  decreasing  the  parameters  of  the  shell,  seen  for 
the  most  part  in  the  groups  from  the  temperate  latitudes  of  both  hemispheres 
(the  Punctidae  and  the  Helicodiscidae)  together  with  the  retention  of  a 
simple  sexual  apparatus. 

2.  The  complication  of  the  apertural  margin  because  of  the  formation 
of  dental  armature  (the  Polygyridae  and  certain  specialized  tropical  Endo- 
dontidae)  .  The  erection  of  this  taxon  is  not  correlated  vidth  any  complication 
of  the  sexual  apparatus. 

3.  The  gradual  reduction  of  the  shell  with  a  parallel  significant 
complication  of  the  male  and  female  genitalia.  This  tendency  can  be  seen 
in  the  many  Helixina  (the  Gastrodontoidea  and  the  Vitrinoidea) .   In  other 
instances  the  reduction  of  the  shell  is  not  accompanied  with  any  significant 
complication  of  the  genitalia  (most  Zonitinia) . 

There  are  three  small  groups  related  to  the  ancient  Punctoidea  whose 
precise  systematic  position  is  still  problematic.   First  there  is  the  mono- 
typic  family  Thyrophorellidae.   The  sole  genus  of  the  family  with  its  unique 
sinistral  shell  has  the  upper  part  of  the  aperture  strongly  projecting  ante- 
riorly and  this  part  is  semi -moveable  attached  to  the  rest  of  the  shell. 


The  genitalia  cast  no  light  on  the  provenance  of  this  group  because  of  their 
simple  structure;  the  only  feature  which  is  worthy  of  attention  is  the  long, 
slightly  fusiform  vagina  which  is  characteristic  for  many  lower  Endodontinia. 
Boettger  (1962)  suggested  that  this  group  should  be  associated  with  the  Ario- 
phantoidea  (i.e.,  in  our  opinion,  Vitrinoidea) ;  however,  the  complicated 
organization  of  the  sexual  system  of  the  latter  is  characteristic,  and  we, 
therefore,  follow  Thiele  (1931)  and  prefer  to  place  the  Thyrophorellidae 
in  the  Endodontinia,  establishing  for  it  a  separate  superfamily  to  emphasize 
its  uniqueness. 

The  second  group  of  debatable  systematic  placement  is  the  Pleurodisci- 
dae.  At  present  the  family  is  unanimously  placed  with  the  lower  Orthurethra 
(essentially  the  Pupillina)  for  the  reason  that  in  Pleurodiscus  both  urethras 
are  open.  However,  this  feature  should  not  serve  as  a  criterion  for  such 
a  high  taxonomic  level.  The  simplicity  of  the  genitalia  of  Pleurodiscus 
is  primary  although  such  a  reduction  of  the  complicated  characteristics  of 
the  Pupillina,  aside  from  the  specialized  Vertiginidae,  does  not  attain  the 
state  of  terminal  attachment  for  the  sexual  retractor  muscle  as  occurs  in 
Pleurodiscus;  this  feature  is  quite  characteristic  for  many  Endodontinia. 
As  for  the  shell,  it  is  of  the  type  near  the  one  which  is  conceivably  or- 
ginal  for  the  Helixina.  All  the  non-bulimuloid  shells  of  the  Pupillina 
have  a  diameter  not  greater  than  3-4  mm  for  reasons  discussed  above.  At 
the  same  time  the  shell  of  Pleurodiscus  reaches  15  mm  in  width.  Finally 
it  is  not  out  of  place  here  to  put  the  question:   what  in  general  associates 
Pleurodiscus  with  the  Pupillina  aside  from  the  formal  resemblance  in  the 
structure  of  the  renopericardial  complex?  One  can  only  answer  this  question: 
nothing . 


From  these  reasons  we  consider  the  family  to  be  one  of  the  lowest  mem- 
bers of  the  suborder  Helixina  which  retained  the  orthurethran  type  of  ex- 
cretory apparatus . 

A  third  familial  group  questionably  included  in  the  Endodontinia  is 
the  Systrophiidae.   The  shell  has  a  typical  endodontid  appearance.   The 
sexual  system  lacks  appendages  and  an  epiphallus  is  not  marked.  Character- 
istic is  the  terminal  attachment  of  the  sexual  retractor  muscle;  in  other 
words,  the  Systrophiidae  are  typical  members  of  Endodontinia.   Nevertheless 
they  are  always  associated  with  the  typically  predatory  families  (the  Zoniti- 
dae  and  the  Trigonochlamydidae) .   However,  the  predatory  specialization  takes 
place  in  various  taxa  of  the  Geophila,  and  one  can  observe  its  independent 
origin  in  several  lineages.   The  Endodontinia  constitute  no  exception. 
Naturally  the  change  to  the  predatory  condition  is  accompanied  by  correspond- 
ing changes  in  the  structure  of  the  radula,  foot,  and  renopericardial 

Solem  (1975)  has  convincingly  shown  that  the  Helicodiscidae  should  be 
an  independent  family. 

Among  the  Helixinia  the  only  obligatory  predators  are  the  Rhytidoidea 
reflected  also  in  their  outer  appearance.  Further,  this  is  also  one  of  the 
rare  cases  when  the  predator  has  a  non-aulacopod  foot.  Nevertheless  the  char- 
acter of  the  mobility  of  these  animals  deserves  a  thorough  investigation, 
(cf.  supra.  Fig.  3,  VI-VII). 

In  Fig.  7  it  is  seen  that  the  Euconulidae  and  the  Gastrodontidae  are 
removed  from  the  Zonitidae  and  associated  as  the  Gastrodontoidea  with  the 
Vitrinoidea.   Their  removal  from  the  Zonitidae  was  explained  earlier  (Schileyko, 
1972a).  There  are  no  serious  objections  to  the  placement  of  these  families 


near  the  Vitrinoidea  (Van  Mol  and  Van  Bruggen,  1971;  Verdcourt,  1960). 
Moreover,  the  association  of  the  Helicarionidae  with  the  Vitrinoidea  is 
confirmed  by  the  fact  that  in  the  Vitrinid  genus  Semilimax  there  is  a  sarco- 
belum  of  characteristic  structure  (Hubendick,  1953  and  our  own  data). 

The  organization  and  the  phylogeny  of  the  higher  Helixinia  --Helicoi- 
dea  of  authors  --  has  already  been  considered  (Shileyko,  1978).  Therefore,  - 
we  will  confine  ourselves  to  the  main  points  here.   Basically  it  must  be 
emphasized  that  it  is  impossible  to  associate  the  Helicoidea  and  the  Hygromioi- 
dea  to  one  another  on  anything  but  superfamilial  grounds.  Thus,  if  we 
were  to  take  the  most  archaic  groups  among  these  taxa,   namely  the  Humboldtiani- 
dae  and  Trichiinae,  we  would  discover  two  distinctly  different  methods  of 
the  placement  of  the  stylophores  (Vide  supra) .  There  is  another  sharp  dif- 
ference between  these  superf  ami  lies.'   in  the  Helicoidea,  the  reservoir  of 
the  spermatheca  is  connected  by  bodies  of  connective  tissue  to  the  bottom 
of  the  mantle  cavity  and  presses  into  the  base  of  the  albumen  gland:   in 
the  Hygromioidea  the  spermatheca  is  not  bound  up  with  the  mantle  cavity 
but  is  joined  to  the  spermoviduct . 

The  Helicodontoidea  apparently  is  a  heterogenous  group  and  at  pre- 
sent it  is  difficult  to  determine  its  phylogenetic  connections.   One  may 
only"  note  that  there  are  four  types  of  sexual  systems  (Helicondonta,  Lind- 
holmiola,  Oestophora,  and  Mastigophallus)  whose  phyletic  succession  cannot 
be  traced.   The  shell  alone  unites  them  in  one  superfamily  --  flat,  multi- 
whorled,  with  reflected  apertural  margins. 

The  organization  of  the  Sphincterochilidae  has  been  discussed  often 
(Schileyko,  1972b;   Forcart,  1972a,  b) ;  the  unique  structure  of  the  foot 
and  the  sexual  apparatus  has  been  emphasized  as  reflecting  a  considerable 


distinction  of  the  family  from  other  helicoid  groups.   At  the  same  time  the 
association  of  the  Sphincterochilidae  with  the  Helicoidea  is  not  in  doubt. 
This  is  due  most  of  all  to  the  typically  helicoid  shell  of  Sphincterochili- 
dae which  only  too  clearly  relates  the  group  to  the  Helicoidea.  Nevertheless, 
if  one  were  to  disassociate  oneself  from  the  prejudice  exerted  by  external 
appearance  and  compare  the  sexual  system  of  the  Sphincterochilidae  with  the 
genitalia  of  the  Vitrinoidea,  it  is  not  difficult  to  see  how  the  Sphinctero- 
chilidae naturally  fits  into  a  scheme  more  closely  related  to  vitrinoids 
than  to  the  helicoids;  the  sarcobelum  is  armed  with  a  mucous  gland;  the 
penis  has  a  sac-like  appendage  and  a  flagellum  (cf.  Solem,  1960;  Van  Mol, 
1968,  1970,  1973).  The  low  degree  of  specialization  of  the  sexual  appara- 
tus in  the  Sphincterochilidae  forces  one  to  see  it  as  a  group  close  to 
the  ancestral  form  of  the  Vitrinoidea  though  still  noticeably  divergent. 
In  the  Vitrinoidea,  the  tendency  to  shell  reduction  is  characteristic.  The 
higher  members  of  the  superfamily  are  already  slugs  or  semi-slugs.   The  lower 
Vitrinoidea  have  well  developed  shells,  and  the  ancestors  of  Sphinctero- 
chilidae, possessing  a  normal  shell,  adapted  to  living  in  arid  conditions; 
their  development  was  directed  toward  the  thickening  of  the  shell  walls  and 
the  formation  of  other  adaptive  features  (i.e.  white  color  and  sharply  nar- 
rowed aperture) . 

The  infraorder  Zonitinia  derived  independently  from  the  Helixina 
from  ancient  Endodontinia  and  are  clearly  separable  into  two  superf amilies : 
1)  Zonitoidea,  in  which  the  process  of  shell  loss  can  be  observed  as  the 
lineage  turns  predatory;  and  2)  the  Parraacelloidea  which  completely  or  parti- 
ally loses  the  shell  in  going  from  herbivorous  to  omnivorous. 

The  peculiarities  of  the  organization  of  the  suborder  Limaxina, 


composed  exclusively  of  shell-less  forms,  has  been  discussed  by  Likharev 
5  Victor  (1979). 

In  summary,  in  examining  the  evolutionary  paths  in  the  suborders 
and  infraorders,  one  may  note  the  following  lines  of  specialization  and  de- 
velopment of  different  organs  and  structures  which  appeared  independently 
and  convergently  in  all  taxa, 

1.  The  appearance  of  sigmurethry  with  complete  closing  of  the  urethra. 

2.  The  formation  of  the  aulacopod  foot  often  accompanied  by  the  ap- 
pearance of  a  tripartite  sole. 

3.  The  reduction  of  the  shell  and  the  evolution  of  the  slug  form. 

4.  The  transition  to  a  predatory  state;  this  process  was  often  associ- 
ated with  the  elaboration  of  a  higher  direction  of  specialization. 

Acknowledged  is  the  criticism  and  help  given  the  author  by  N.  N.  Akramovski, 
Ya.  I.  Starobogatov  and  I.M.  Likharev. 


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Schileyko,  A.  A.   19T6a.   The  paths  of  evolution  of  the  excretory 

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Schileyko,  A.  A.   l9T6b.   Features  of  the  organization  and  systematics 

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Schileyko,  A.  A.   1978.   Mollusks  of  the  superfamily  Helicoidea  of  the 
fauna  of  USSR.   Fauna  USSR.,  new  series,  no.  117,  mollusks  3, 
sect.  6:  I-38I+. 

Schileyko,  L.  V.  and  A.  A.  Schileyko.   1975-   On  the  morphology  of  the 

proximal  section  of  the  sexual  system  of  Stylommatophora  (Gastropoda, 
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reports  of  the  higher  educational  school  for  biological  sciences), 
no.  1.   pp.  7-13- 


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Fig.  1  Comparative  shell  volume  occupied  by  the  molluscan 
body  (darkened)  in  regard  to  the  extent  of  the  umbilicus.   The 
shells  have  the  same  proportions  and  the  same  contours. 
A.  Widely  umbilicate  B.  narrovly  umbilicate. 

Fig.  2  The  types  of  placement  of  the  grooves  in  the  foot  of 
the  Geophila. 

I)  from  under  the  buccal  flap  extend  a  few  short  suprapedal 
grooves   (Partula) 

II)  the  same  type   (Placostylus,  Helicophanta,  Urocoptidae) 

III)  only  peripedal  grooves   ( Cerion,  Oleacina,  Paryphanta, 
Helix,  Subulinidae,  Clausiliidae) 

IV)  suprapedal  and  peripedal  grooves  (sic)  (the  author  means 
with  one  pair  of  pedal  grooves  and  one  pair  of  suprapedal 
grooves   (Nesovitrea  and  Cecilioides ) 

V)  two  pairs  of  suprapedal  and  one  pair  of  pedal  grooves. 
(Macrochlamys,  Vitrina,  Oxychilus ,  Trigonochlamys ,  Limax) 

VI)  one  central  pedal  groove   (Rhytida) 

VII)  one  central  pedal  groove  and  a  pair  of  suprapedal 
grooves   (Zonitoides ) 

VIII )  a  pair  of  suprapedal  grooves   (Vitrea) 

IX )  two  pairs  of  suprapedal  grooves   (Discus ,  Arion) 

X)  groove  absent,  but  waves  of  locomotor  contractions  pass 
only  along  the  central  part  of  the  sole   (Achatina) 

XI)  hypothetical  variant  of  the  appearance  of  pedal  grooves 

at  the  expense  of  the  differentiation  of  the  central  field. 

Incisions  in  II-XI  made  on  the  level  indicated  by  dashed  line  in  I. 


Figure   1 

Figure   2 

Fig.  3.   Two  methods  of  flagellum  formation 

I-IV)  The  descent  of  the  seminal  duct  (vas  deferens)  distally. 

V-VIII)  The  junction  of  the  lower  end  of  the  seminal  duct  (vas- 
deferens)  with  the  apical  part  of  the  epiphallus. 

Fig.  h.    Schematic  representation  of  the  male  sexual  section  of  Pupillina. 
A1-A5  -  parts  of  the  penial  appendix 
PR  -  sexual  or  penial  retractor  muscle 
F  -  flagellum 
C  -  caecum 


Figure   3 

Figure   h 

Fig.  5-  External  view  and  form  of  the  sperraatophore  in  Pseudonapaeus 
albipliciatus  (Mts.)   (Pupillina,  Buliminidae)  above  —  a  sperraatophore 
taken  from  the  spermatheca;  below  —  the  epiphallus  and  cross  section 
through  the  "tail"  of  the  sperraatophore.   The  walls  of  the  sperraatophore 
shown  in  sections. 

Fig.  6.  The  reproductive  system  and  its  structural  details  in 
different  Achatinina. 

A  -  Placostylus  shongi  (Less.)  from  New  Zealand   (Bulimulidae) ; 

the  reproductive  system  and  with  penis  opened 

B  -  Antidrymaeus  inusitatus  (Fult.)  from  Brazil   (Orthalicidae)  ; 
(Orthalicidae  not  separated  from  Bulimulidae)  the  reproductive 
system  and  the  partially  opened  spermatheca 

V  -  the  same  with  the  penis  opened 

G  -  the  same  showing  the  organs  of  the  mantle  cavity 

D  -  Helicophanta  magnifica  (Per.)  from  Madigascar   (Acavidae);  the 
reproductive  apparatus  and  the  opened  penis 

E  -  Rumina  decollata  (L.)  from  Algeria   (Subulinidae) ;  with 
the  reproductive  apparatus  and  the  opend  penis 

r  \.- 


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•  op  iji/e^f  uj 





3cp:  jjsoiuf 






3Dp^j  mftiSity 


















































































































































Fig.  8.   Partula  otaheitana  (Brug. )  from  Tahiti,  Tahita-nua. 
A  -  external  view  of  the  soft  parts  taken  from  the  shell 
B  -  organs  of  the  mantle  cavity 
V  -  reproductive  system 

G  -  part  of  the  herraaphoroditic  duct  and  quadrivia  or  ferti- 
lization chamber 
D  -  penis  opened 
SU  -  secondary  urethra 
RV  -  respiratory  vein 
K  -  kidney 
R  -  rectum 
RS  -  rectal  sinus 

Fig.  9.   Cerion  glans  (Kust.)  from  Cuba. 
A  -  reproductive  apparatus 
B  -  quadrivia  or  fertilization  chamber 
V  -  atrial  section  of  the  genitalia 

G  -  roof  of  the  mantle  cavity  and  cross  section  of  the  kidney 
PA  -  penial  appendix 
AV  -  atrial  valve 
AM  -  atrial  muscle 

DS  -  diverticula  of  the  spermatheca 
RV  -  respiratory  vein 
K  -  kidney 

PU  -  primary  urethra 
R  -  rectum 
S  -  spermatheca 

Fig.  10.   Oleacina  solidula  (Pfr.)  from  Cuba.   Left  -  reproductive 
system  and  cross  sections  through  uterus  and  vas  deferens.   Center  -  penis 
in  the  region  of  its  appendages,  opened.   Right  -  organs  of  the  mantle  cavity, 

DPA  -  distal  penial  appendage 

PPA  -  proximal  penial  appendage 

SU  -  secondary  urethra 

K  -  kidney 

PU  -  primary  urethra 

RO  -  renal  orifice  (orifice  of  primary  urethra) 

S  -  spermatheca 

E  -  epiphallus 



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